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Improving Seed Germination of the Eggplant Rootstock Solanum Torvum by Testing Multiple

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Improving Seed Germination of the Eggplant Rootstock Solanum Torvum by Testing Multiple 1 Improving seed germination of the eggplant rootstock Solanum torvum by testing multiple 2 factors using an orthogonal array design 3 4 R.H.G. Ranila, H.M.L. Niranb, M. Plazasc, R.M. Fonsekaa, H.H. Fonsekad, S. Vilanovac, I. Andújarc, 5 P. Gramazioc, A. Fitac, J. Prohensc,* 6 7 aDepartment of Crop Science, Faculty of Agriculture, University of Peradeniya, Old Galaha Road, 8 Peradeniya 20400, Sri Lanka 9 bAgriculture Research Station, Girandurukotte 90750, Sri Lanka 10 cInstituto de Conservacion y Mejora de la Agrodiversidad Valenciana, Universitat Politècnica de 11 València, Camino de Vera 14, 46022 Valencia, Spain 12 dHorticultural Crop Research and Development Institute, Gannoruwa Road, Peradeniya 20400, Sri 13 Lanka 14 15 *Corresponding author. Tel: +34 963879424; fax: +34 963879422. 16 E-mail addresses: [email protected] (R.H.G. Ranil), [email protected] (H.M.L. Niran), 17 [email protected] (M. Plazas), [email protected] (R.M. Fonseka), 18 [email protected] (H.H. Fonseka), [email protected] (S. Vilanova), 19 [email protected] (I. Andújar), [email protected] (P. Gramazio), [email protected] (A. Fita), 20 [email protected] (J. Prohens) 21 22 1 23 ABSTRACT 24 Solanum torvum is a highly vigorous relative of eggplant that is resistant to a number of harmful 25 soil-borne diseases and is compatible for grafting with eggplant. Being a potential rootstock, this 26 plant frequently presents poor and erratic germination, which makes its practical use difficult. We 27 used an L8 (27) orthogonal array design to evaluate the primary effects of seven factors (soaking of 28 seeds, scarification with sodium hypochlorite (NaClO), application of gibberellic acid (GA3), use of 29 potassium mitrate (KNO3) as a moistening agent, cold stratification, application of a heat shock, and 30 light irradiation during germination) at two levels (L0 and L1) using four germination parameters 31 (early and final germination, germination rate and vigour index) in fresh S. torvum seeds. Solanum 32 torvum seeds had a strong dormancy with no germination in the untreated seeds and high early and 33 final germination (approximately 100%) in certain treatments. An evaluation of the main effects 34 revealed highly positive effects on germination from seed soaking, and the use of GA3, KNO3, and 35 light irradiation, whereas NaClO scarification had a negative effect. The application of cold 36 stratification and heat shock treatments also had a positive effect on seed germination but to a lesser 37 extent than the other treatments. An improved proposed protocol that consisted of subjecting seeds 38 to soaking, the application of GA3 and KNO3, cold stratification, heat shock, and light irradiation 39 was validated and demonstrated to be highly effective, with seed germination success greater than 40 60% being observed at 3 d and final germination reaching a plateau at 6 d. A second validation 41 experiment using a commercial growing substrate also showed a high emergence (approximately 42 50%) at 7 d and a final germination of approximately 80% was recorded with application of the 43 improved protocol. The seed germination protocol that we have developed will facilitate the use of 44 S. torvum as a rootstock for eggplant and its use in breeding programmes. Our results also reveal 45 that orthogonal array designs are a powerful tool for establishing improved protocols for seed 46 germination. 47 48 Keywords: Solanum torvum, seed, germination, dormancy, orthogonal arrays, main effects 2 49 50 1. Introduction 51 52 Solanum torvum Sw., commonly known as turkey berry, devil’s fig or pea eggplant, is a 53 wild bush of neotropical origin that belongs to the “spiny Solanum” (subgenus Leptostemonum) 54 group (Levin et al., 2006). This species has become naturalised and is sometimes invasive in 55 tropical areas of Africa, Asia, and Australia; also, this species is occasionally cultivated, primarily 56 in Southeast Asia and Africa, for its edible fruits (Gousset et al., 2005; Nyadanu and Lowor, 2015). 57 Solanum torvum is of great interest as a rootstock for eggplant (S. melongena L.), as the plant is 58 highly vigorous, fully graft-compatible with eggplant scions (Gisbert et al., 2011b; Moncada et al., 59 2013), and possesses resistance to a wide range of soil pathogens, such as Verticillium dahliae, 60 Ralstonia solanacearum, Fusarium oxysporum, and root-knot-nematodes (Bletsos et al., 2003; 61 Gousset et al., 2005; Bagnaresi et al., 2013), as well as being tolerant to abiotic stresses (Schwarz et 62 al., 2010). Furthermore, eggplant fruits produced on scions grafted onto S. torvum are of good 63 quality (Gisbert et al., 2011b; Moncada et al., 2013; Miceli et al., 2014). Additionally, grafting 64 eggplant on S. torvum reduces translocation of the heavy metal cadmium (Cd) from the roots to the 65 aerial part (Arao et al., 2008) and may minimize the negative effects on the fruit quality from Cd 66 soil contamination (Savvas et al., 2010). Because of these desirable traits, S. torvum also represents 67 a genetic resource of strong relevance to the introgression breeding of eggplant (Kumchai et al., 68 2013) 69 The primary limitation for the practical use of S. torvum as a rootstock in the commercial 70 production of grafted eggplant plants, as well as in breeding programmes, is the poor, irregular and 71 erratic germination due to dormancy in seeds (Ginoux and Laterrot, 1991; Miura et al., 1993; 72 Gousset et al., 2005; Hayati et al., 2005). This characteristic has even led to the proposed use of 73 vegetative propagation to overcome the seed germination problem (Miceli et al., 2014). The 74 breaking of dormancy, which is a common phenomenon among wild Solanum species (Taab and 3 75 Andersson, 2009; Wei et al., 2010; Kandari et al., 2011; Tellier et al., 2011), and the enhancement 76 of germination can be achieved using combinations of many different physical (e.g., seed soaking, 77 manual scarification, cold stratification, heat shocks, light irradiation, and magnetic fields) and/or 78 chemical (e.g., scarification with acidic or basic chemicals, plant growth regulators, and osmotic 79 treatments) treatments (Finch-Savage and Leubner-Metzger, 2006; Bewley et al., 2013; Holubowicz 80 et al., 2014). 81 Determining the critical combination of factors that permit the enhancement of germination 82 in S. torvum seeds is important to develop improved protocols for seed germination in this species 83 for its use as a rootstock and for breeding purposes (Hayati et al., 2005; Gisbert et al., 2011a). The 84 influence of several potentially key factors affecting a variable, in this case seed germination, can 85 be determined by studying one factor at a time (as achieved by Hayati et al., 2005). However, this 86 process greatly reduces efficiency when there is an interdependency of factors or when it is 87 impractical to isolate and test each variable individually. Full factorial designs, which are much 88 more efficient in determining the optimal combination of factors, may require large and costly 89 experiments when many factors are involved (Onyiah, 2008; Rao et al., 2008). An alternative 90 commonly used in industrial applications are orthogonal (Taguchi) arrays (Roy, 2010), which allow 91 the main effects of a large number of factors to be estimated with a limited number of treatments. 92 Although orthogonal arrays have been successfully applied to address the problem of determining 93 adequate combinations of factors in biological and biotechnological processes (Rao et al., 2008; 94 Assemi et al., 2012; Sedghi et al., 2014; Vasilev et al., 2014), their use for establishing seed 95 germination protocols has been highly limited (Wu et al., 2011; Poinapen et al., 2013) and largely 96 overlooked. 97 In this work, we evaluate the primary effects of seven factors potentially involved in the 98 release from dormancy and enhancement of seed germination in dormant seeds of S. torvum using 99 an orthogonal array experimental design. The improved protocol established according to the results 100 was then tested and validated. The results from this work will provide information for improving 4 101 the seed germination of S. torvum. These findings will also contribute to facilitating its use as a 102 rootstock and as a source of variation in breeding programmes. At the same time, this work aims to 103 demonstrate the potential of orthogonal array designs for establishing efficient seed germination 104 protocols. 105 106 2. Materials and Methods 107 108 2.1. Seed materials and germination conditions 109 110 Fresh seeds of S. torvum accession No. 55953 (originally purchased from Sunshine Seeds, 111 Ahlen, Germany) were extracted from physiologically ripe fruits of plants cultivated in an open 112 field at Universitat Politècnica de València (Valencia, Spain). Five-month-old seeds of S. 113 melongena accession No. BBS-188/B (landrace from Ivory Coast), with high germination values 114 (>90%), were also used as a control for the validation of the improved treatment developed for the 115 germination of S. torvum seeds. 116 Depending on the experiment, seeds were germinated in Petri dishes (9.0 × 2.5 cm; Phoenix 117 Biomedical, Mississauga, Ontario, Canada) on a layer of 0.5 cm of embedded hydrophilic cotton 118 covered by filter paper or sown at a depth of 7 mm in plastic pots (9 × 9 × 9.5 cm) containing 119 commercial nursery growing substrate (Neuhaus Huminsubstrat N3, Lassmann-Dellmann, Geeste, 120 Germany). Twenty-five evenly distributed seeds were placed in each Petri dish or pot. Seeds were 121 sown in Petri dishes and pots at the beginning of the experiments (day 0) in a climatic chamber with 122 a 14-h light / 10-h dark photoperiod at 25ºC constant temperature. A light irradiance of 600 123 mmol·m-2·s-1 was provided by GRO-LUX F36W/GRO (Sylvania, Danvers, MA, USA) fluorescent 124 tubes.
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